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Hamiota, a New Genus of Freshwater Mussel (Bivalvia: Unionidae) from the Gulf of Mexico Drainages of the Southeastern United States Kevin J

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Hamiota, a New Genus of Freshwater Mussel (Bivalvia: Unionidae) from the Gulf of Mexico Drainages of the Southeastern United States Kevin J Iowa State University From the SelectedWorks of Kevin J. Roe 2005 Hamiota, a new genus of freshwater mussel (Bivalvia: Unionidae) from the Gulf of Mexico drainages of the southeastern United States Kevin J. Roe Paul D. Hartfield Available at: https://works.bepress.com/kevin_roe/9/ THE NAUTILUS 119(1):1–10, 2005 Page 1 Hamiota, a new genus of freshwater mussel (Bivalvia: Unionidae) from the Gulf of Mexico drainages of the southeastern United States Kevin J. Roe Paul D. Hartfield Delaware Museum of Natural History U.S. Fish and Wildlife Service 4840 Kennett Pike 6578 Dogwood View Parkway, Suite A Wilmington, DE 19807 USA Jackson, MS 39213 USA [email protected] Paul࿞Hartfi[email protected] ABSTRACT MATERIALS AND METHODS Hamiota, a new genus of freshwater mussel containing four A list of specimens examined is included in Appendix 1. species formerly assigned to the genus Lampsilis Rafinesque, Acronyms used in the text are: Academy of Natural Sci- 1820, is described. In addition to the genus Lampsilis, mem- ences of Philadelphia (ANSP), Delaware Museum of bers of Hamiota had previously been placed in the genera Vil- Natural History (DMNH), Florida Museum of Natural losa Frierson, 1927, and Ligumia Swainson, 1840. Several char- History (UF), United States National Museum (USNM), acters including the packaging of their larvae in a superconglu- University of Alabama Unionid Collection (UAUC), and tinate lure to attract host fishes, placement and shape of the Mississippi Museum of Natural Science (MMNS). Ab- marsupia, and release of glochidia through the excurrent si- breviated synonymies are presented for each taxon and phon, support the recognition of these species as a distinct include novel combinations and publications with illus- genus. tration. Measurements were taken to the nearest 0.05 mm using dial calipers. SYSTEMATICS INTRODUCTION Family Unionidae Rafinesque, 1820 Following the discovery in 1988 by Robert Butler of the Tribe Lampsilini von Ihering, 1901 first superconglutinate lure ensnared on a snag in a trib- Hamiota new genus. Type species: Hamiota subangulata (Lea, utary of the Choctawhatchee River, a number of publi- 1840) by original designation cations (e.g., Haag et al., 1995; Hartfield and Butler, 1997; O’Brien and Brim Box, 1999; Blalock-Herod et al., Diagnosis: A monophyletic group of freshwater bi- 2002) have confirmed through direct observation the valves (Roe et al., 2001) in which all of the glochidia are supposition that these unique lures are produced by four released simultaneously encased in mucous packages species of freshwater mussels endemic to the Gulf of that are referred to as superconglutinates (Haag et al., Mexico drainages of the southeastern United States. 1995, fig. 1). The superconglutinate lure exits the mantle Herein, we confirm earlier published suggestions that cavity via the excurrent opening and is encased within a these four species represent a distinct genus of fresh- transparent mucous tube (Hartfield and Butler, 1997; water mussels (Fuller and Bereza, 1973; O’Brien and O’Brien and Brim Box, 1999). When acted upon by wa- Brim Box, 1999). The recognition of this genus is based ter currents the superconglutinate mimics the move- on several characters including the production of the su- ments of a swimming fish, and has been shown to elicit perconglutinate lure, and the unique shape and place- attacks from fishes (Haag and Warren, 1999). The mar- ment of the marsupia (the region of the demibranchs supium is restricted to the ventral portion of the outer where female unionoid mussels brood developing lar- demibranchs of female mussels (Figure 1). The precise vae), and is supported by molecular evidence (Roe et al., shape and pigmentation of the marsupia, as well as the 2001). Use of marsupial features is consistent with pre- degree of posterior mantle margin development, varies vious designations of unionid ‘‘divisions’’ initiated by across species. Simpson (1900a) and continued by Heard and Guckert Description: Members of this genus are small- to me- (1970). The designation of Hamiota increases the num- dium-sized freshwater bivalves, and adult valves gener- ber of North American unionids genera to 50 (Turgeon ally are between 45–100 mm in length. Shells range et al., 1998). from ovate to elliptical in outline, and are somewhat Page 2 THE NAUTILUS, Vol. 119, No. 1 Figures 1, 2. Inner mantle and outer demibranchs of gravid superconglutinate-producing mussels in the genus Hamiota and inner mantle and outer demibranchs typical of non-superconglutinate-producing mussels of the genera Lampsilis, Ligumia, and Villosa. 1. Female Hamiota australis. 2. Female Villosa vibex. compressed to moderately inflated. Shell thickness rang- and colors may include purple, red, black, or white. In es from heavy to thin. Sexes display some degree of di- females, the mantle margins anterior to the branchial morphism in shell shape. Shells of male mussels are typ- opening are elaborated to varying degrees. The mor- ically more acutely pointed posteriorly, whereas shells of phology of the glochidial valves of members of Hamiota female mussels display an expanded posterior margin. is similar to that of members of Lampsilis or Villosa Periostracum is typically smooth, but can be very glossy (O’Brien and Brim Box, 1999). in some species. Background color ranges from dark- Etymology: Hamiota ϭ angler. Derived from the brown and black through chestnut-brown to straw-yel- Greek word hamus, meaning hook. This name refers to low. Black to bright green rays of variable width are of- the means by which members of this genus attract host ten present and may be limited to the posterior slope, fishes by packaging their parasitic larvae in a lure that or cover the entire disk. Nacre color is typically white mimics a small fish. although other colors such as salmon or blue may be seen as well, particularly in the beak cavities or the pos- Remarks: Species of Hamiota generally have been terior margins. The marsupia are often asymmetrical in treated as Lampsilis due to the similarity in shape and shape; the anterior portion is typically broadest, tapering coloration of their shells. Some authors also have in- toward the posterior end. The ventral margin of the mar- cluded these species in the genus Villosa or Ligumia, supium is darkly pigmented in gravid females. Pigmen- due to shell shape, thickness, and/or ornamentation of tation of marsupia varies across species and populations the mantle flap. The shells of Hamiota species are in- K. J. Roe and P. D. Hartfield, 2005 Page 3 deed similar to the shells of these genera, and the man- mantle flap is typically not expanded and is reddish in tle flap may exhibit characters of both Lampsilis and color. Marsupia of H. altilis are finely tapered at each Villosa. However, in Hamiota, the marsupia is restricted end when immature, becoming broadly rounded on the to the ventral half of the posterior portion of the outer ends in most populations, tapering anteriorly in others. demibranchs, while in Lampsilis, Ligumia, and Villosa Marsupia color is a dark reddish-brown or black along the marsupium fully occupies the water tubes of the pos- the margin and white above. The anus is usually pig- terior portion of the outer demibranchs (Figure 1). Un- mented red and the incurrent and excurrent siphons are like members of Lampsilis, which release larvae through reddish-brown to black. The glochidia of H. altilis are pores in the ventral edge of the demibranchs, members described by Haag et al. (1999). of Hamiota release the superconglutinate lure through Type Material: Unio altilis Conrad, 1834, Lectotype the excurrent opening. The placement and shape of the ANSP 56419 (Figures 3, 4) here designated. Type lo- marsupium and the extraordinary method of glochidial cality: Alabama River, near Claiborne [Monroe Co., Al- release and host fish attraction, the superconglutinate, abama]. are uniquely derived characters in the Lampsilini that Unio clarkianus Lea, 1852, Type not found. Type local- warrant genus-level recognition. ity: Williamsport, [Maury Co.], Tennessee; Georgia or Alabama. Hamiota altilis (Conrad, 1834) new combination Unio doliaris Lea, 1865, Lectotype USNM 84936, here designated. Type locality: Etowah River, Georgia. Unio altilis Conrad, 1834; Conrad, 1834: 43, pl. 2, fig. 1; Unio gerhardtii Lea, 1862, Holotype USNM 25711 by Chenu, 1845: 21, pl. 1, fig. 1; Reeve, 1865: pl. 23, fig. 109. monotypy. Type locality: Chattanooga, Georgia. Margarita (Unio) altilis (Conrad, 1834); Lea, 1836: 24. Margaron (Unio) altilis (Conrad, 1834); Lea, 1852a: 27. Remarks: The most variable species included in Ham- Lampsilis altilis (Conrad, 1834); Simpson, 1900a: 529; Par- iota, H. altilis, is endemic to the Mobile River Basin. malee and Bogan, 1998: 125, pl. 47. Some of the conchological variation is undoubtedly eco- Unio clarkianus Lea, 1852; Lea, 1852b: 251; Lea, 1852c: 273, phenotypic in nature, although the extent and nature of pl. 21, fig. 30; Lea, 1852d: 29, pl. 21, fig. 30. Margaron (Unio) clarkianus (Lea, 1852); Lea, 1852a: 27. the variation in shell shape and pigmentation has not Lampsilis clarkianus (Lea, 1852); Simpson, 1900a: 532. been adequately explored. Unio gerhardtii Lea, 1862; Lea, 1862a: 168; Lea, 1862b: 208, Conrad (1834) in his original description did not iden- pl. 31, fig. 277; Lea, 1862c: 30, pl. 31, fig. 277. tify a primary type. Johnson and Baker (1973) identified Margaron (Unio) gerhardtii (Lea, 1862); Lea, 1870: 35. ANSP 56419 as the figured holotype, although the spec- Lampsilis (Lampsilis) gerhardtii (Lea, 1862); Simpson, 1900a: imen label indicates the locality as ‘‘Ogeechee R., Ga.’’ 532. Johnson and Baker (1973) state that the label is in error, Unio doliaris Lea, 1865; Lea, 1865: 88; Lea, 1868: 260, pl. 32, and ‘‘probably was mixed with ANSP 46418, which is fig. 75; Lea, 1869: 20, pl. 32, fig. 75. labeled ‘Claiborne, Alabama’, by error.’’ Conrad (1834) Margaron (Unio) doliaris (Lea, 1865); Lea, 1870: 42.
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